Lacunal allocation and gas transport capacity in the salt marsh grass Spartina alterniflora

Summary Lacunal allocation as the fraction of the total cross sectional area of leaves, stem bases, rhizomes, and roots was determined in both tall and short growth forms of Spartina alterniflora collected from natural monospecific stands. The results indicate that in both growth forms lacunal allocation is greater in stem bases and rhizomes than in leaves and roots and that tall form plants allocate more of their stem and rhizome to lacunae than short form plants.Measurements made in natural stands of Spartina alterniflora suggest that total lacunal area of the stem base increases with increasing stem diameter and that stem diameter increases with increasing plant height and above-ground biomass. However, the fraction of cross section allocated to lacunae was relatively constant and increased only with the formation of a central lacuna.Experimental manipulations of surface and subsurface water exchange were carried out to test the influence of flooding regime on aerenchyma formation. No significant differences in lacunal allocation were detected between plants grown in flooded (reduced) and drained (oxidized) sediments in either laboratory or field experiments. While aerenchyma formation in Spartina alterniflora may be an adaptation to soil waterlogging/anoxia, our results suggest that lacunal formation is maximized as a normal part of development with allocation constrained structurally by the size of plants in highly organic New England and Mid-Atlantic marshes.The cross sectional area of aerenchyma for gas transport was found to be related to the growth of Spartina alterniflora with stands of short form Spartina alterniflora exhibiting a lower specific gas transport capacity (lacunal area per unit below ground biomass) than tall form plants despite having a similar below-ground biomass supported by a 10 fold higher culm density. The increased specific gas transport capacity in tall vs. short plants may provide a new mechanism to explain the better aeration, higher nutrient uptake rates and lower frequency of anaerobic respiration in roots of tall vs. short Spartina alterniflora.     

Estimation of primary production using five different methods in a Spartina alterniflora salt marsh

The aboveground production of Spartina alterniflora in a salt marsh in Barataria Bay, Louisiana, USA was estimated using five different harvest methods: peak standing crop (PSC), Milner-Hughes, Smalley, Wiegert-Evans, and Lomnicki et al., and a non-destructive method based on measurement of stem density and longevity. Annual production estimates were 831 ± 41, 831 ± 62, 1231 ± 252, 1873 ± 147 and 1437 ± 96 g dry wt m^–2 for each method, respectively. The average longevity of individually tagged young shoots was 5.2 ± 0.2 months, equivalent to an annual turnover rate of 2.3 crops per year. Among the five methods, Wiegert-Evans and Lomnicki et al. were considered more accurate than the other three because they corrected for mortality losses between sampling times. The Lomnicki et al. method was preferred over the Wiegert-Evans method because of its greater simplicity.     

Oxygen loss from Spartina alterniflora and its relationship to salt marsh oxygen balance

Spartina alterniflora has been reported to lose significant amounts of oxygen to its rhizosphere with potentially important effects on salt-marsh biogeochemical cycling and plant productivity. The potential significance of this oxidative pathway was evaluated using laboratory split-chamber experiments to quantify oxygen loss from intact root systems under a wide variety of pre-treatment and incubation conditions including antibiotics to inhibit microbial respiration. The aerenchyma system of S. alterniflora was found to transport O₂, N₂, Ar, and CH₄ from above-ground sources to its below-ground roots and rhizomes. While non-respiratory gases were observed to move from the lacunae to water bathing the root systems, net O₂ loss did not occur; instead oxygen present outside of the roots/rhizomes was consumed. Net oxygen loss was found when resistance to gas transport was reduced in the lacunae-rhizosphere pathway by placing the root systems in a gas phase and when plant respiration was significantly reduced. Root system respiration appeared to be the major variable in the plant oxygen balance. When root and rhizome respiration was inhibited using poisons or lowered by cooling, the oxygen deficit was greatly reduced and oxygen loss was indicated. The effect of seasonal temperature changes on root system oxygen deficit presents a possible explanation as to why Spartina produces root systems with respiration rates that cannot be supported by gas transport. Overall, while oxygen loss from individual plant roots is likely, integrating measured root system oxygen loss with geochemical data indicates that the mass amount of oxygen lost from S. alterniflora root systems is small compared to the total oxygen balance of vegetated salt marsh sediments.     

Spread of Exotic Cordgrasses and Hybrids (Spartina sp.) in the Tidal Marshes of San Francisco Bay, California, USA

Four species of exotic cordgrass (Spartina sp.) occur in the San Francisco estuary in addition to the California native Spartina foliosa. Our goal was to map the location and extent of all non-native Spartina in the estuary. Hybrids of S. alterniflora and S. foliosa are by far the most numerous exotic and are spreading rapidly. Radiating from sites of deliberate introduction, S. alterniflora and hybrids now cover ca. 190 ha, mainly in the South and Central Bay. Estimates of rate of aerial increase range from a constant value to an accelerating rate of increase. This could be due to the proliferation of hybrid clones capable of rapid expansion and having superior seed set and siring abilities. The total coverage of 195 ha by hybrids and other exotic cordgrass species is slightly less than 1% of the Bay's tidal mudflats and marshes. Spartina anglica has not spread beyond its original 1970s introduction site. Spartina densiflora has spread to cover over 5 ha at 3 sites in the Central Bay. Spartina patens has expanded from 2 plants in 1970 to 42 plants at one site in Suisun Bay. Spartina seed floats on the tide, giving it the potential to export this invasion throughout the San Francisco estuary, and to estuaries outside of the Golden Gate. We found isolated plants of S. alterniflora and S. densiflora in outer coast estuaries north of the Bay suggesting the likelihood for the San Francisco Bay populations to found others on the Pacific coast.     

Restoration of a Spartina alterniflorasalt marsh following a fuel oil spill,New York City,NY

In the seven years since 2.5 million liters of No. 2fuel oil spilled into the Arthur Kill, the Salt MarshRestoration Team of New York City Parks has restoredSpartina alterniflora(Salt Marsh Cordgrass)to 2.43 ha of shoreline, planting seedlings andtransplants in areas of low salt marsh severelyimpacted by oil. Restoration was undertaken to haltsurface erosion and the loss of surviving vegetationand remnant peat. Biomass, stem density, flowerdensity, height of plants, rhizome spread, basal areaand plant cover were monitored for a minimum of threeyears between 1993 and 1997. Unplanted SeverelyImpacted (USI) reference sites and Existing Vegetation(EV) in the severe impact zone were also monitored.Ninety-five percent of the surface area denuded ofvegetation by oil at USI reference sites remainedunvegetated seven years after the spill. SedimentTotal Petroleum Hydrocarbon (TPH) content was measuredbefore and after plantings and these results indicateda high level of persistent contamination. Residual oilwas not the determining factor in the survival or lossof Spartina alternifloraseedlings andtransplants at planting sites. Instead, plantspacing, shoreline morphology, wave energy generatedby passing vessels, and predation by Brantacanadensis(Canada Goose) were the most criticalfactors. Three years after planting, the abovegroundbiomass at two of the three restoration sites wascomparable to the biomass at existing and restoredeastern North American salt marshes. Seedlings spaced30 cm apart provided rapid cover at two of three sitesand contributed to overall survival. Where initialsurvival was high, goose predation was low. Whereinitial survival was low, shoreline morphology andwave energy were the primary causes of mortality, withgoose predation an important secondary cause. Bedformsat one planting site indicated a high energyshoreline as did sediment texture results for allsites. The fetch along the Arthur Kill is not longenough to generate such high-wave energy, but passingvessels do generate plunging and surging breakers ona regular basis. A reduction in the frequency of oilspills since 1990 has been beneficial to the saltmarshes along the Arthur Kill. A reduction in otheranthropogenic stresses would further enhance theplanted and existing vegetation.     

Nitrogen addition does not reduce belowground competition in a salt marsh clonal plant community in Mississippi (USA)

Previous studies have suggested that belowground competition for nutrients influences plant zonation in salt marshes. In this study, I tested the hypothesis that competition for nitrogen structured a clonal plant community in a nitrogen-limited salt marsh in coastal Mississippi, USA. In contrast to most previous field studies that have investigated mechanisms of competition, I examined clonal growth responses of established genets of a nitrogen-demanding low-intertidal species (Spartina alterniflora) to nitrogen addition and the removal of a nitrogen-conserving high-intertidal species (Juncus roemerianus). Nitrogen addition stimulated clonal invasion of the Juncus zone by Spartina but did not reduce the significant competitive effects of Juncus on Spartina. Simulated Juncus shade did not reduce invasion of the Juncus zone by Spartina, indicating that belowground competition reduced clonal invasion. In the last year of the study, the border shifted unexpectedly towards the Spartina zone, resulting in competitive displacement of Spartina by Juncus. Nitrogen addition did not prevent or slow this displacement, further contradicting the nitrogen competition hypothesis. Although growth rates were much more strongly limited by nitrogen in Spartina than in Juncus, nitrogen addition did not cause the displacement of Juncus by Spartina after three growing seasons. I conclude that zonation of Spartina and Juncus is maintained by preemption of space and greater tolerance of low nitrogen supplies by Juncus in the high marsh. These results contrast sharply with findings of reduced belowground competition with nutrient addition in previous studies and highlight the important role of nutrient-mediated competition for space between clonal plants.     

Fungal biomass and decomposition in Spartina maritima leaves in the Mondego salt marsh (Portugal)

Spartina maritima (Curtis) Fernald is a dominant species in the Mondego salt marsh on the western coast of Portugal, and it plays a significant role in estuarine productivity. In this work, leaf litter production dynamics and fungal importance for leaf decomposition processes in Spartina maritima were studied. Leaf fall was highly seasonal, being significantly higher during dry months. It ranged from 42 g m^-2 in June to less than 6 g m^-2 during the winter. Fungal biomass, measured as ergosterol content, did not differ significantly between standing-decaying leaves and naturally detached leaves. Fungal biomass increased in wet months, with a maximum of 614 g g^-1 of ergosterol in January in standing-decaying leaves, and 1077 g g^-1 in December, in naturally detached leaves, decreasing greatly in summer. Seasonal pattern of fungal colonization was similar in leaves placed in litterbags on the marsh-sediment surface. However, ergosterol concentrations associated with standing-decaying and naturally detached leaves were always much higher than in litterbagged leaves, suggesting that fungal activity was more important before leaf fall. Dry mass of litterbagged leaves declined rapidly after 1 month (about 50%), mostly due to leaching of soluble organic compounds. After 13 months, Spartina leaves had lost 88% of their original dry weight. The decomposition rate constant (k) for Spartina maritima leaves was 0.151 month^-1.     

A technique for distinguishing between bacterial and non-bacterial respiration in decomposing Spartina alterniflora

A number of antibiotics were used to suppress bacterial activity in decomposing Spartina alterniflora. The effectiveness of each treatment was quantified using INT formazan vital staining and epifluorescent microscopy. Bacterial suppression of selected treatments was verified using standard plate count procedures. Chloramphenicol treated samples (exhibiting 87–90% bacterial suppression) were analyzed respirometrically and found to consume only 30% less O₂ than controls. Non-bacterial respiration (probably fungal) accounted for 70% of the respiration.     

Sediment addition enhances transpiration and growth of Spartina alterniflora in deteriorating Louisiana Gulf Coast salt marshes

Transpiration, leaf conductance, net photosynthesis, leaf growth, above-ground biomass and regeneration of new culms were studied in a rapidly subsiding Spartina alterniflora Lois. salt marsh following the addition at 47 and 94 Kg m^–2 of new sediment. Plant growth was enhanced in response to sediment addition as was evident by a significant increase in leaf area, above-ground biomass production and regeneration of new culms (p 0.05). Leaf conductance and transpiration rates were significantly greater in sediment treated plants than in control plants (p 0.05). Enhanced production of culms per unit area of marsh resulted in increased leaf area which allowed a greater capacity for net photosynthesis and contributed to increases in above-ground biomass of sediment treated plots.     

Diel methane emissions in stands of Spartina alterniflora and Suaeda salsa from a coastal salt marsh

In this study, we aimed to understand the influence of plant type on the monthly variations of diel CH₄ fluxes from Spartina alterniflora and Suaeda salsa of coastal salt marshes at three growth stages (July, August and September). Dissolved CH₄ concentrations in porewater and sediment redox potentials were monitored, as were aboveground plant biomass and stem densities. CH₄ fluxes exhibited clear monthly variations and peaked in September in the S. alterniflora and S. salsa mesocosms. However, no discernible diel variation was observed in the CH₄ flux in the S. salsa mesocosm, probably due to its weak gas transport capacity. By contrast, notable diel variations of CH₄ flux with the peak of 1.42 and 3.67 mg CH₄ m⁻² h⁻¹ at 12:00 and the lowest of 0.75 and 2.11 mg CH₄ m⁻² h⁻¹ at 3:00 or 6:00 were observed in the S. alterniflora mesocosm on 11 August and 11 September, respectively, but not in July mainly due to low plant biomass masking diel variations in the porewater CH₄ concentration. The ratios of the maximum flux to minimum flux over the course of the day in the S. alterniflora mesocosm on 10 July, 11 August and 11 September were 1.28, 1.89 and 1.76, respectively, and corresponding values for porewater CH₄ concentration were 1.31, 1.39 and 1.17, respectively. CH₄ flux significantly correlated with CH₄ concentration in porewater, and both were significantly related to air temperature. These findings indicate that CH₄ production and CH₄ flux at the middle growth stage (August) exhibited greater responses to changes in air temperature, which in turn induced the higher diel variation. The higher diel cycle for CH₄ flux in August than in September was likely due to the higher proportion of CH₄ oxidized during diffusion within the aerenchyma system. Our results suggest that the extent of diel variations in CH₄ flux may have depended on the gas transport capacity of plants, and the highest diel variation occurred at the middle growth stage.     

First record of the sea anemone Diadumene lineata (Verrill 1871) associated to Spartina alterniflora roots and stems, in marshes at the Bahia Blanca estuary, Argentina

We report the occurrence of the orange-striped green anemone Diadumene lineata (Verrill 1871) (=Haliplanella lineata) in salt marshes at the Bahía Blanca Estuary for the first time in August 2005. We also found this species attached to roots and stems of Spartina alterniflora, an association that has never been registered before. After their determination, sampling was performed during a year to evaluate seasonal abundance of this sea anemone. Results showed that D. lineata was present through the whole year, indicating the existence of a stable population. All individuals sampled were found attached to roots or stems of S. alterniflora, with the higher abundances detected in summer. Further studies are necessary to precise the potential effects of this exotic sea anemone on salt marsh communities.     

Specific criteria in Grania (Oligochaeta,Enchytraeidae)

The structure of the penial bulb and male efferent duct system of Grania species may be used in addition to setal pattern and spermathecal shape to distinguish species. Six penial bulb types are distinguished: (1) a simple, small, glandular bulb surrounding the male pore; (2) a small, glandular bulb, with a large, associated, dorso-medial gland mass; (3) a small glandular bulb, medial to the male pore, with an elongate male bursa (the aglandular sac), the vas deferens exitting directly into the invaginated male pore; (4) a glandular bulb with an aglandular sac and a small, cuticular stylet embedded in the bulb, extending from the ectal end of the vas deferens; (5) a glandular bulb and an aglandular sac with a long stylet extending from the vas deferens, through the bulb into the sac; and (6) glandular bulb reduced or absent, with or without an aglandular sac; with a long stylet and other prominent modifications, usually muscular, of the vas deferens. The details of the male duct structure were consistent within specimens grouped on the basis of setal distribution and shape and detailed spermathecal structure. Diverse male duct patterns are found within the polytypic species G. macrochaeta and G. postclitellochaeta. The positions of the spermathecal and male pores in their respective segments are distinctive for some species.     

互花米草生物量变化对盐沼沉积物有机碳的影响

以江苏王港典型互花米草(Spartina alterniflora)盐沼湿地为研究对象,分析光滩及互花米草滩沉积物中有机碳的水平和垂向分布特征,了解互花米草生物量的季节动态变化,探讨二者之间的相互关系,在此基础上研究互花米草生物量分布和季节变化对沉积物中有机碳(TOC)含量的影响。结果表明,互花米草枯落物中的有机碳数量在两个月内衰减了40%,而表层沉积物中TOC含量及其中互花米草来源TOC所占比例的变化,均与互花米草地表枯落物量的季节变化存在两个月的"相位差",这与枯落物快速分解时间大致吻合,说明枯落物是表层沉积物中TOC的重要来源。高达60%的互花米草地下生物量分布在0—20cm深度内,该深度范围内沉积物中TOC含量较高,且TOC主要来源于互花米草。此外,不同深度TOC含量与地下生物量之间存在良好的正相关关系,说明地下生物量是影响沉积物TOC含量的重要因子。研究区互花米草年固碳能力为2274g m-2a-1,盐沼沉积物中TOC埋藏速率达到了470 g m-2a-1,是地表一个重要的碳汇;同时研究区每年向近岸水域输出大量的TOC,是近岸海域生态系统的一个重要碳源。     

Decomposition processes of Spartina maritima in a salt marsh of the Basque Country

Decomposition dynamics of aerial parts and root-rhizomes of Spartina maritima in a Basque Country salt marsh was studied, using litter bags placed on the soil surface and buried 10 cm below ground. Aerial parts of the plant in aboveground position showed higher breakdown rates than samples placed belowground. There was no significant difference found between aerial parts and root-rhizomes buried. Nitrogen and phosphorus followed different dynamics (seasonal changes and progressive losses) that may be a consequence of distinctive mineralization pathways. The low faunal richness and densities belowground reflect the unfavourable life conditions in such a situation and, to a certain extent, the lower decomposition rates of buried litter. Four mathematical expressions that fit the data are presented and discussed.     

Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats

A common but often less tested explanation for the successful invasion of alien species is that invasive alien species outcompete their co-occurring natives, which may not always be the case. In this study, we established artificial environmental gradients in a series of pot experiments with controlled environments to investigate the effects of salinity, sediment type and waterlogging on the performance of and interactions between Phragmites australis (native) and Spartina alterniflora (alien), which generally co-exist in the saline intertidal zones of Chinese and American coasts. Significant effects of salinity and waterlogging were detected on biomass production and morphological characteristics of S. alterniflora and P. australis, and the competitive interactions between the two species were found to vary with all three environmental factors in our experiments. Relative Neighbor Effect (RNE) analyses indicate that competitive dominance of S. alterniflora occurred under the conditions of high salinity, sandy sediment and full immersion, whereas P. australis showed competitive dominance under the conditions of low salinity and non-immersion. Our results suggest that S. alterniflora might outcompete P. australis under conditions present in early salt marsh succession, which support the viewpoint that the outcomes of competition between co-occurring native and invasive alien plants depend on the growing conditions. The implication of this study is that in response to the environmental changes expected from seawater intrusion and sea-level rise, the range of S. alterniflora is expected to expand further in the Yangtze River estuary in the future.     

Decomposition of Spartina anglica,Elytrigia pungens and Halimione portulacoides in a Dutch salt marsh in association with faunal and habitat influences

Decomposition of Spartina anglica, Elytrigia pungens and Halimione portulacoides was studied for 20.5 months in situ in two habitats on a salt marsh in The SW Netherlands. Litter bags of three different mesh sizes were used to exclude meio- and/or macrofauna. The middle-marsh habitat was flooded more frequently than the plant-debris habitat in the highest marsh zone. Decomposition of the three species followed an exponential pattern of decay: instantaneous decay rates varied from 0.0026 to 0.0054 per day. Decay rates were significantly influenced by habitat factors and fauna, while there was a significant interaction between plant species and habitat. In case of a significant meio- and/or macrofauna effect, this became noticeable 12–16 weeks after the start of decomposition and resulted in a difference of 5–10% ash-free dry weight remaining after 20.5 months. Nematodes were the dominant microfaunal group in the plant litter. Densities were influenced by habitat conditions but not by resource quality, season and meio- and/or macrofauna. Only initial C/N and C/P ratios were correlated with differences in decomposition rates between the plant species. During the later stages of decomposition N and P concentrations of the plant litter were higher in the plant-debris habitat than in the middle-marsh habitat, probably as a result of fluctuating detritivores densities. The course of the decomposition process differed per plant species and per habitat. The results of this study underline the importance of knowledge of long-term decomposition rates.     

Waterlogging responses in dune,swale and marsh populations of Spartina patens under field conditions

Summary Soil waterlogging responses were examined in three Spartina patens populations along a steep flooding gradient in coastal Louisiana. Root anatomy and physiological indicators of anaerobic metabolism were examined to identify and compare flooding responses in dune, swale and marsh populations, while soil physicochemical factors were measured to characterize the three habitats. Soil waterlogging increased along the gradient from dune to marsh habitats and was accompanied by increases in root porosity (aerenchyma). Aerenchyma in marsh roots was apparently insufficient to provide enough oxygen for aerobic respiratory demand, as indicated by high root alcohol dehydrogenase activities and low energy charge ratios. Patterns of root metabolic indicators suggest that dune and swale roots generally respired aerobically, while anaerobic metabolism was important in marsh roots. However, in each population, relatively greater soil waterloging was accompanied by differences in enzyme activities leading to malate accumulation. In dune and swale roots under these circumstances, depressed adenylate energy charge ratios may have been the result of an absence of increased ethanol fermentation. These trends suggest that: 1) Aerenchyma formation was an important, albeit incomplete, long-term adaptation to the prevalent degree of soil waterlogging. 2) All populations adjusted root metabolism in response to a relative (short-term) increase in soil waterlogging.     

Habitat size, flora, and fauna: Interactions in a tidal saltwater marsh

Anthropogenic habitat fragmentation is increasingly problematic in both terrestrial and aquatic systems. Fragmentation reduces the size of habitat patches, so examining the effect of patch size on community structure can provide insight into the potential effects of fragmentation. In this study, we examined the effect of habitat size on the density of Spartina alterniflora shoots in tidal saltwater marshes, as well as on the two predominant macrofaunal species, the marsh periwinkle Littoraria irrorata and fiddler crabs Uca spp. We estimated the density of shoots in three different marsh habitats, (1) large island marshes, (2) small island marshes, and (3) large fringing marshes, in Indian Field Creek, York River, Chesapeake Bay. We manipulated shoot density in each of the marsh types to distinguish between the effects of marsh grass density and marsh type on crab and Littoraria densities in the system. We found significant differences in grass density among the three marsh types as well as significant species-specific effects of grass density, marsh type, and distance from edge on faunal abundance. Decreasing the shoot density resulted in a decrease in Littoraria density in the large marshes. Littoraria density increased with distance from edge in the small marshes and in the first 5 m of the fringing marshes, then decreased with distance from edge after 5 m in the fringing marshes. Shoot density had a negative effect on crabs in both the large and small marshes. These results suggest that fragmentation would have a negative effect on the community structure by lowering the densities of both the flora and fauna.     

Mobility of metals in salt marsh sediments colonised by <Emphasis Type="Italic">Spartina maritima</Emphasis> (Tagus estuary,Portugal)

Chemical associations of Zn, Pb, Cu, Co and Cd were determined using a sequential extraction procedure in sediments colonised by S. maritima in three salt marshes within the Tagus estuary: Rosário, Corroios and Pancas. Concentrations of these metals were also analysed in above- and belowground parts of Spartina maritima, as well as in sediments colonised by the plant. The highest metal concentrations in sediments were found in the marshes near the industrial and urban areas, whereas metal concentrations in plants were not significantly different among sites. This was thought to be a consequence of differences observed in metal bioavailability: Metals in Pancas, the least polluted location, were largely associated to easily accessible fractions for plant uptake, probably as a result of low organic matter content and high sandy fraction in sediments. S. maritima was able to induce the concentration of metals between its roots in the three salt marshes. The results obtained in this study indicate that S. maritima could be useful to induce phytostabilisation of metals in sediments, although the effectiveness to modify chemical associations is highly dependent on existing sediment parameters, and thus different results could be obtained depending on site characteristics. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

Biomass production,photosynthesis, and leaf water relations of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> under moderate water stress

The perennial smooth cordgrass, Spartina alterniflora, has been successfully introduced in salty ecosystems for revegetation or agricultural use. However, it remains unclear whether it can be introduced in arid ecosystems. The aim of this study was to investigate the physiological response of this species to water deficiency in a climate-controlled greenhouse. The experiment consisted of two levels of irrigation modes, 100 and 50% field capacities (FC). Although growth, photosynthesis, and stomatal conductance of plants with 50% FC were reduced at 90 days from the start of the experiment, all of the plants survived. The water-stressed plants exhibited osmotic adjustment and an increase in the maximum elastic modulus that is assumed to be effective to enhance the driving force for water extraction from the soil with small leaf water loss. An increase in the water use efficiency was also found in the water-stressed plants, which could contribute to the maintenance of leaf water status under drought conditions. It can be concluded that S. alterniflora has the capacity to maintain leaf water status and thus survive in arid environment.